No Effect of Male Courtship Intensity on Female Remating in the Butterfly <Emphasis Type="Italic">Pieris napi</Emphasis>

In Pieris napi, female fitness increases with number of matings, but wild females mate at an unexpectedly low rate. From a sexual conflict perspective this could be because males manipulate female remating, or alternatively, because wild females experience costs associated with remating which are not applicable under laboratory conditions. To get an indication which sex controls remating and/or the different sexes’ relative costs and benefits of remating, we here test whether female mating frequency is affected by male courtship intensity. We found no effect on female mating frequency or lifespan. This indicates that (i) females control remating and their optimal mating frequency is lower compared to males, or (ii) males can manipulate female remating. We argue that both these alternatives may apply simultaneously to P. napiand that they are inseparable.     

Why Do Female Callosobruchus maculatus Kick Their Mates?

Sexual conflict is now recognised as an important driver of sexual trait evolution. However, due to their variable outcomes and effects on other fitness components, the detection of sexual conflicts on individual traits can be complicated. This difficulty is exemplified in the beetle Callosobruchus maculatus, where longer matings increase the size of nutritious ejaculates but simultaneously reduce female future receptivity. While previous studies show that females gain direct benefits from extended mating duration, females show conspicuous copulatory kicking behaviour, apparently to dislodge mating males prematurely. We explore the potential for sexual conflict by comparing several fitness components and remating propensity in pairs of full sibling females where each female mated with a male from an unrelated pair of full sibling males. For one female, matings were terminated at the onset of kicking, whereas the other’s matings remained uninterrupted. While fecundity (number of eggs) was similar between treatments, uninterrupted matings enhanced adult offspring numbers and fractionally also longevity. However, females whose matings were interrupted at the onset of kicking exhibited an increased propensity to remate. Since polyandry can benefit female fitness in this species, we argue that kicking, rather than being maladaptive, may indicate that females prefer remating over increased ejaculate size. It may thus be difficult to assess the presence of sexual conflict over contested traits such as mating duration when females face a trade off between direct benefits gained from one mating and indirect benefits from additional matings.     

The benefits of male ejaculate sex peptide transfer in Drosophila melanogaster

The accessory gland protein (Acp) ejaculate molecules of male Drosophila melanogaster mediate sexual selection and sexual conflict at the molecular level. However, to date no studies have comprehensively measured the timing and magnitude of fitness benefits to males of transferring specific Acps. This is an important omission because without this information it is not possible to fully understand the strength and form of selection acting on adaptations such as Acps. Here, we measured the fitness benefits to males of ejaculate sex peptide (SP) transfer. SP is of interest because it is a candidate for mediating sexual conflict: its frequent receipt reduces female fitness. In single matings with virgin females SP is known to increase egg laying and decrease receptivity. Hence, we predicted that SP could: (i) boost a male’s absolute paternity by increasing offspring production and delaying female remating and/or (ii) boost relative paternity share. We tested these predictions using two different lines of SP‐lacking males, in both two‐mating and free‐mating assay conditions. SP transfer conferred higher absolute, but not relative, male reproductive success. In matings with virgin females, SP transfer increased mating productivity and delayed remating and hence the onset of sperm competition. In already mated females, SP transfer did not elevate absolute progeny production, but did increase intermating intervals and hence the period over which a male could gain paternity. Consistent with this, under free‐mating conditions over an extended period, we detected a ‘per‐mating’ fitness benefit for males transferring SP. These benefits are consistent with a role for SP in mediating conflict, with SP acting to maximize short‐term fitness benefits for males.     

Mating-induced inhibition of remating in female Mediterranean fruit flies Ceratitis capitata

Mechanisms producing inhibition of remating in mated female Mediterranean fruit flies Ceratitis capitata, were investigated by matings with surgically altered males. Comparison of remating by females mated with either intact control males or males with a shortened penis, showed that ejaculate or a physical stimulus of penis insertion caused remating inhibition for at least 10 days after first mating. Remating frequency at two days after mating was significantly higher in females mated to castrated (spermless) males than in females mated to sham-operated control males. This difference disappeared by day four after mating, indicating that sperm cause a shorter-term inhibition of remating than does a normal first mating. Other factors in addition to sperm must therefore play a role in inhibition of remating.     

Costly sexual harassment in a beetle

Abstract The optimal number of mating partners for females rarely coincides with that for males, leading to sexual conflict over mating frequency. In the bruchid beetle Callosobruchus maculatus, the fitness consequences to females of engaging in multiple copulations are complex, with studies demonstrating both costs and benefits to multiple mating. However, females kept continuously with males have a lower lifetime egg production compared with females mated only once and then isolated from males. This reduction in fitness may be a result of damage caused by male genitalia, which bear spines that puncture the female’s reproductive tract, and/or toxic elements in the ejaculate. However, male harassment rather than costs of matings themselves could also explain the results. In the present study, the fitness costs of male harassment for female C. maculatus are estimated. The natural refractory period of females immediately after their first mating is used to separate the cost of harassment from the cost of mating. Male harassment results in females laying fewer eggs and this results in a tendency to produce fewer offspring. The results are discussed in the context of mate choice and sexual selection.     

Sexual conflict and cooperation under naturally occurring male enforced monogamy

An evolutionary conflict often exists between the sexes in regard to female mating patterns. Females can benefit from polyandry, whereas males mating with polyandrous females lose reproductive opportunities because of sperm competition. Where this conflict occurs, the evolution of mechanisms whereby males can control female remating, often at a fitness cost to the female, are expected to evolve. The fitness cost to the female will be increased in systems where a few high status males monopolise mating opportunities and thus have limited sperm supplies. Here we show that in the cockroach Nauphoeta cinerea, a species where males enforce female monogamy in the first reproductive cycle, males that have become sperm depleted continue to be able to manipulate female remating behaviour. Although the manipulation severely decreases fecundity in females mated to sperm-depleted males, males benefit, increasing their relative fitness by preventing other males from reproducing. Our results suggest that there is selection on maintaining the mechanism of manipulation rather than maintaining sperm numbers. Taken with previous research on sexual conflict in N. cinerea, this study suggests that the causes and consequences of sexual conflict are complex and can change across the life history of an individual.     

The Effects of Experimentally Induced Polyandry on Female Reproduction in a Monandrous Mating System

Females of most insect species maximize their fitness by mating more than once. Yet, some taxa are monandrous and there are two distinct scenarios for the maintenance of monandry. While males should always benefit from inducing permanent non‐receptivity to further mating in their mate, this is not necessarily true for females. Since females benefit from remating in many species, cases of monandry may reflect successful male manipulation of female remating (i.e. sexual conflict). Alternatively, monandry may favor both mates, if females maximize their fitness by mating only once in their life. These two hypotheses for the maintenance of monandry make contrasting predictions with regards to the effects of remating on female fitness. Here, we present an experimental test of the above hypotheses, using the monandrous housefly (Musca domestica) as a model system. Our results showed that accessory seminal fluid substances that males transfer to females during copulation have a dual effect: they trigger female non‐receptivity but also seem to have a nutritional effect that could potentially enhance female fitness. These results suggest that monandry is maintained in house flies despite potential benefits that females would gain by mating multiply.     

Receptive females mitigate costs of sexual conflict

Males typically gain fitness from multiple mating, whereas females often lose fitness from numerous mating, potentially leading to sexual conflict over mating. This conflict is expected to favour the evolution of female resistance to mating. However, females may incur male harassment if they refuse to copulate; thus, greater female resistance may increase costs imposed by males. Here, I show that the evolution of resistance to mating raises fitness disadvantages of interacting with males when mating is harmful in female adzuki bean beetles, Callosobruchus chinensis. Females that were artificially selected for higher and lower remating propensity evolved to accept and resist remating, respectively. Compared with females that evolved to accept remating, females that evolved to resist it suffered higher fitness costs from continuous exposure to males. The costs of a single mating measured by the effect on longevity did not differ among selection line females. This study indicates that receptive rather than resistant females mitigate the fitness loss resulting from sexual conflict, suggesting that even though mating is harmful, females can evolve to accept additional mating.     

Remating responses are consistent with male postcopulatory manipulation but not reinforcement in D. pseudoobscura

Reinforcement occurs when hybridization between closely related lineages produces low‐fitness offspring, prompting selection for elevated reproductive isolation specifically in areas of sympatry. Both premating and postmating prezygotic behaviors have been shown to be the target of reinforcing selection, but it remains unclear whether remating behaviors experience reinforcement, although they can also influence offspring identity and limit formation of hybrids. Here, we evaluated evidence for reinforcing selection on remating behaviors in Drosophila pseudoobscura, by comparing remating traits in females from populations historically allopatric and sympatric with Drosophila persimilis. We found that the propensity to remate was not higher in sympatric females, compared to allopatric females, regardless of whether the first mated male was heterospecific or conspecific. Moreover, remating behavior did not contribute to interspecific reproductive isolation among any population; that is, females showed no higher propensity to remate following a heterospecific first mating than following a conspecific first mating. Instead, we found that females are less likely to remate after initial matings with unfamiliar males, regardless of species identity. This is consistent with one scenario of postmating sexual conflict in which females are poorly defended against postcopulatory manipulation by males with whom they have not coevolved. Our results are generally inconsistent with reinforcement on remating traits and suggest that this behavior might be more strongly shaped by the consequences of local antagonistic male–female interactions than interactions with heterospecifics.     

Females of the Cellar Spider Discriminate Against Previous Mates

Mate choice for novel partners should evolve when remating with males of varying genetic quality provides females with fitness‐enhancing benefits. We investigated sequential mate choice for same or novel mating partners in females of the cellar spider Pholcus phalangioides (Pholcidae) to understand what drives female remating in this system. Pholcus phalangioides females are moderately polyandrous and show reluctance to remating, but double‐mated females benefit from a higher oviposition probability compared to single‐mated females. We exposed mated females to either their former (same male) or a novel mating partner and assessed mating success together with courtship and copulatory behaviours in both sexes. We found clear evidence for mate discrimination: females experienced three‐fold higher remating probabilities with novel males, being more often aggressive towards former males and accepting novel males faster in the second than in the first mating trial. The preference for novel males suggests that remating is driven by benefits derived from multiple partners. The low remating rates and the strong last male sperm precedence in this system suggest that mating with novel partners that represent alternative genotypes may be a means for selecting against a former mate of lower quality.     

同时而非先后一雌多雄交配增加不同温度下食蚜瓢虫的适合度（英文）

【目的】尽管一雌多雄在瓢虫科中常见,但各研究中获得的数据不足以解释雌虫多次交配和一雌多雄的一般适应性意义或适合度后果。本研究以温度为胁迫因子,旨在评价一雌多雄的某些益处（如增加的适合度）是否可传递给后代。【方法】本研究检测了黄斑盘瓢虫Coelophora saucia (Mulsant) 3种交配处理中的适合度：一雌一雄（与同一雄虫交配5次,1次/d）,先后一雌多雄（与5头不同的雄虫依次交配5次,即每天与新的雄虫交配1次）,以及同时一雌多雄（放进5头雄虫,任由雌虫选择雄虫,交配5次,1次/d）。观察了各交配处理不同温度下（25, 27和 30℃）繁殖力、卵的育性、后代发育和存活。【结果】结果表明,经历一雌多雄然后进行交配选择或竞争的雌性的繁殖能力最强,后代能在更广温度范围内最好地适应发育和存活。但先后一雌多雄交配的雌性与一雌一雄交配的雌性的繁殖能力相似。【结论】结果说明,在无交配选择或雄性竞争的条件下,一雌多雄的益处不明显。这可能是由于在依次射精的雄性间存在精子竞争,或由于雌性的隐性选择。据我们所知,本研究中观察发现的无交配选择时不表现一雌多雄的益处的现象,之前在昆虫中未观察到过。     

Theoretical influence of female mating status and remating propensity on male sperm allocation patterns

Theoretical models predict that males should allocate more sperm in matings where the immediate risk of sperm competition is high. It has therefore often been argued that males should invest less sperm in matings with virgin females compared with matings with already mated females. However, with relatively polyandrous females, high sperm competition risk will covary with high sperm competition intensity leading to more unpredictable conditions, as high competition intensity should favour smaller ejaculates. With the use of a genetic algorithm, we found that males should allocate more sperm in matings with virgin females when female mating frequency is relatively high, whereas low remating rates will select for higher effort in matings with nonvirgin females. At higher remating rates, first male sperm precedence favours larger ejaculates in matings with virgin females and second male precedence favours the reverse. These results shed some light on several findings that have been difficult to explain adaptively by the hitherto developed theory on sperm allocation.     

Fitness components of male and female winter moths (Operophtera brumata L.) (Lepidoptera, Geometridae) relative to measures of body size and asymmetry

In this article we present data from two experiments on theassociation between individual asymmetry and fitness in thewinter moth. We performed a mate selection experiment and comparedasymmetry and body size of mated and unmated males collectedin the field. Individual asymmetry was not associated with copulationprobability, adult life span, or body size, even though body sizeis a reliable indicator of larval and pupal survival, femalefecundity, adult life span, and thus expected fitness. Therewas only a weak positive effect of body size on mating success,contrary to the strong effect of female size on male choicefound in previous experiments. Both males and females were capableof repeated mating, and the number of matings was correlatedwith female size, but neither with male body size nor with adultasymmetry. Yet, females engaged in repeated matings more frequentlyif they were first mated to a more asymmetrical male. This mayindicate that more asymmetrical males lose paternity due tofemale remating, although direct paternity analyses need tobe carried out. In addition, repeated mating may be uncommonunder field situations. In conclusion, the relationship betweenindividual asymmetry and fitness seems to be at best weak inthe winter moth.     

Selection for multiple mating in females due to mates that reduce female fitness

If females are unable to discriminate among males before mating,remating by females that store sperm may have evolved as a hedgeagainst having only "costly" mates (less preferred males thatreduce her fitness). However, the benefit of remating is notguaranteed because she can also mate by chance with anothercostly male. We devised a model to explain the evolution offemale remating by representing female fitness as a functionof the proportion of costly mates. We examined the effect ofa linear, a concave-up, and a concave-down fitness functionand found that only the latter favors the evolution of femaleremating. With a concave-down function, females mating with50% costly mates have nearly the same fitness as do femaleswith none. A biological interpretation for a concave-down functionis that sperm from good males are better at competing with spermfrom costly males or are more preferred by females. A concave-upfunction implies the reverse, whereas a linear function willoccur when sperm are equally competitive. We review specificsituations in nature that might produce a concave-down functionand find evidence that sterility and intragenomic conflict aretwo phenomena capable of driving the evolution of female rematingby our model.     

Sex peptide causes mating costs in female Drosophila melanogaster

Conflicts between females and males over reproductive decisions are common . In Drosophila, as in many other organisms, there is often a conflict over how often to mate. The mating frequency that maximizes male reproductive success is higher than that which maximizes female reproductive success . In addition, frequent mating reduces female lifespan and reproductive success , a cost that is mediated by male ejaculate accessory gland proteins (Acps) . We demonstrate here that a single Acp, the sex peptide (SP or Acp70A), which decreases female receptivity and stimulates egg production in the first matings of virgin females , is a major contributor to Acp-mediated mating costs in females. Females continuously exposed to SP-deficient males (which produce no detectable SP ) had significantly higher fitness and higher lifetime reproductive success than control females. Hence, rather than benefiting both sexes, receipt of SP decreases female fitness, making SP the first identified gene that is likely to play a central role in sexual conflict.     

Nuptial gift consumption influences female remating in a scorpionfly: male or female control of mating rate?

In the scorpionfly Panorpa cognata, males provide females with saliva secretions as nuptial food gifts. Consequently, females derive material benefits and possibly also genetic benefits from multiple matings. Females therefore generally should have a high motivation to remate. Males, on the other hand, do not share this interest, which will generate a sexual conflict over remating interval, possibly leading to male adaptations that prevent females from remating with other males. In this study, I found that mated females were less prone to copulate than virgin females, despite female benefits of multiple matings. Further, I found that the remating interval was significantly longer if the first copulation was long compared to shorter matings. This effect does not entirely depend on copulation duration per se, but on the amount of saliva, that a female is consuming during copulation. These results suggest a mating-induced refractory period and can be interpreted as male manipulation of female remating behaviour mediated through substances in the nuptial gift. Alternatively, receiving large nuptial gifts may decrease the prospective direct fitness benefits from further copulations, and thus change optimal female remating rate. Furthermore, gift size has been shown to correlate with male nutritional condition, which may be an indicator of male genetic quality. Females may therefore benefit indirectly by not remating following copulations involving large saliva gifts. In this scenario, female remating interval would be an effect of cryptic female choice.     

Does a lack of mating opportunities explain monandry in the green‐veined white butterfly (Pieris napi)?

In the nuptial gift-giving butterfly, Pieris napi, multiple mating corresponds to higher lifetime fecundity than monandry does. Yet, female mating frequency, which is genetically determined and heritable, varies from strict monandry to a high degree of polyandry. Polyandrous females are known to suffer from reduced longevity when denied the opportunity to mate according to their intrinsic mating frequency. Here we test if monandry is likely to be maintained due to a lack of mating opportunities in the wild. We also explore longitudinal variation in female mating frequency (i.e. lifetime number of matings) and remating rate (i.e. time interval between successive matings). The latter was explored by comparing the mating patterns of females with varying origins in a laboratory, and the former by comparing the mating frequencies of wild and laboratory females. We were able to reveal spatial variation in female mating frequency, with monandry being a clearly more common mating tactic in the north than in the south. However, we did not find direct evidence that either the maintenance of monandry or spatial variation in female mating frequency is promoted by a lack of mating opportunities. Still, our findings may have profound implications, since decreasing mating frequency towards the north generates spatial variation in the strength of sexual selection.     

Visual acuity in larval zebrafish: behavior and histology

Background

Mating plugs that males place onto the female genital tract are generally assumed to prevent remating with other males. Mating plugs are usually explained as a consequence of male-male competition in multiply mating species. Here, we investigated whether mating plugs also have collateral effects on female fitness. These effects are negative when plugging reduces female mating rate below an optimum. However, plugging may also be positive when plugging prevents excessive forced mating and keeps mating rate closer to a females' optimum. Here, we studied these consequences in the gonochoristic nematode Caenorhabditis remanei. We employed a new CO₂-sedation technique to interrupt matings before or after the production of a plug. We then measured mating rate, attractiveness and offspring number.

Results

The presence of a mating plug did not affect mating rate or attractiveness to roving males. Instead, females with mating plugs produced more offspring than females without copulatory plugs.

Conclusions

Our experiment suggests that plugging might have evolved under male-male competition but represents a poor protection against competing males in our experiment. Even if plugging does not reduce mating rate, our results indicate that females may benefit from being plugged in a different sense than remating prevention.     

The evolution of polyandry: multiple mating and female fitness in insects

Theory suggests that male fitness generally increases steadily with mating rate, while one or a few matings are sufficient for females to maximize their reproductive success. Contrary to these predictions, however, females of the majority of insects mate multiply. We performed a meta-analysis of 122 experimental studies addressing the direct effects of multiple mating on female fitness in insects. Our results clearly show that females gain directly from multiple matings in terms of increased lifetime offspring production. Despite a negative effect of remating on female longevity in species without nuptial feeding, the positive effects (increased egg production rate and fertility) more than outweigh this negative effect for moderate mating rates. The average direct net fitness gain of multiple mating was as high as 30-70%. Therefore, the evolutionary maintenance of polyandry in insects can be understood solely in terms of direct effects. However, our results also strongly support the existence of an intermediate optimal female mating rate, beyond which a further elevated mating rate is deleterious. The existence of such optima implies that sexual conflict over the mating rate should be very common in insects, and that sexually antagonistic coevolution plays a key role in the evolution of mating systems and of many reproductive traits. We discuss the origin and maintenance of nuptial feeing in the light of our findings, and suggest that elaborate and nutritional ejaculates may be the result of sexually antagonistic coevolution. Future research should aim at gaining a quantitative understanding of the evolution of female mating rates. Copyright 2000 The Association for the Study of Animal Behaviour.     

Sex Differences in “Time Out” from Reproductive Activity and Sexual Selection in Male Bushcrickets (Orthoptera: Zaprochilinae: <Emphasis Type="Italic">Kawanaphila mirla</Emphasis>)

Animals of many species prefer some partners over others. Discriminating among potential mates causes strong sexual selection that shapes characters and behaviors. In bushcrickets the sexes shows different latencies to remate due to differences in investment in production of the nuptial gift by males and the induced refractory period in females. We conducted experiments with the Australian bushcricket Kawanaphila mirla to test the variation in male mating success by female choice. Male remating intervals under unlimited access to food and mates were around two days, whereas most females did not remate within 12 days. Males had therefore a much shorter “time-out” from mating than females. The adult sex ratio from field samples was near to 1:1. Consequently, the OSR was male-biased with more males than females ready to mate. This male-biased OSR led to mating competition in males and choosiness in females. In a field enclosure with unlimited supply of receptive females the number of matings varied widely between males, with twenty percent of males neglected by the females. The number of matings within this enclosure was neither related to male size nor to song characters, recorded previously in the lab. However, the number of matings by individual males was positively correlated to the size of their spermatophore producing accessory gland. Females appear to prefer males with a large nutritive donation, thereby receiving a direct fitness benefit.